The question of whether sauropod cervicals got longer through ontogeny came up in the comment thread on Mike’s “How horrifying was the neck of Barosaurus?” post, and rather than bury this as a comment, I’m promoting it to a post of its own.

The short answer is, yeah, in most sauropods, and maybe all, the cervical vertebrae did lengthen over ontogeny. This is obvious from looking at the vertebrae of very young (dog-sized) sauropods and comparing them to those of adults. If you want it quantified for two well-known taxa, fortunately that work was published 16 years ago – I ran the numbers for Apatosaurus and Camarasaurus to see if it was plausible for Sauroposeidon to be synonymous with Pleurocoelus, which was a real concern back in the late ’90s (the answer is a resounding ‘no’). From Wedel et al. (2000b: pp. 368-369):

Despite the inadequacies of the type material of Pleurocoelus, and the uncertainties involved with referred material, the genus can be distinguished from Brachiosaurus and Sauroposeidon, even considering ontogenetic variation. The cervical vertebrae of Pleurocoelus are uniformly short, with a maximum EI of only 2.4 in all of the Arundel material (Table 4). For a juvenile cervical of these proportions to develop into an elongate cervical comparable to those of Sauroposeidon, the length of the centrum would have to increase by more than 100% relative to its diameter. Comparisons to taxa whose ontogenetic development can be estimated suggest much more modest increases in length.

Carpenter & McIntosh (1994) described cervical vertebrae from juvenile individuals of Apatosaurus and Camarasaurus. Measurements and proportions of cervical vertebrae from adults and juveniles of each genus are given in Table 4. The vertebrae from juvenile specimens of Apatosaurus have an average EI 2.0. Vertebrae from adult specimens of Apatosaurus excelsus and A. louisae show an average EI of 2.7, with an upper limit of 3.3. If the juvenile vertebrae are typical for Apatosaurus, they suggest that Apatosaurus vertebrae lengthened by 35 to 65% relative to centrum diameter in the course of development.

The vertebrae from juvenile specimens of Camarasaurus have an average EI of 1.8 and a maximum of 2.3. The relatively long-necked Camarasaurus lewisi is represented by a single skeleton, whereas the shorter-necked C. grandis, C. lentus, and C. supremus are each represented by several specimens (McIntosh, Miller, et al. 1996), and it is likely that the juvenile individuals of Camarasaurus belong to one of the latter species. In AMNH 5761, referred to C. supremus, the average EI of the cervical vertebrae is 2.4, with a maximum of 3.5. These ratios represent an increase in length relative to diameter of 30 to 50% over the juvenile Camarasaurus.

If the ontogenetic changes in EI observed in Apatosaurus and Camarasaurus are typical for sauropods, then it is very unlikely that Pleurocoelus could have achieved the distinctive vertebral proportions of either Brachiosaurus or Sauroposeidon.

C6 of Apatosaurus CM 555 – despite having an unfused neural arch and cervical ribs, the centrum proportions are about the same as in an adult.

A few things about this:

From what I’ve seen, the elongation of the individual vertebrae over ontogeny seems to be complete by the time sauropods are 1/2 to 2/3 of adult size. I get this from looking at mid-sized subadults like CM 555 and the hordes of similar individuals at BYU, the Museum of Western Colorado, and other places. So to get to the question posed in the comment thread on Mike’s giant Baro post – from what I’ve seen (anecdata), a giant, Supersaurus-class Barosaurus would not necessarily have a proportionally longer neck than AMNH 6341. It might have a proportionally longer neck, I just haven’t seen anything yet that strongly suggests that. More work needed.

Juvenile sauropod cervicals are not only shorter than those of adults, they also have less complex pneumatic morphology. That was the point of the figure at the top of the post. But that very simple generalization is about all we know so far – this is an area that could use a LOT more work.

I’ve complained before about papers mostly being remember for one thing, even if they say many things. This is the canonical example – no-one ever seems to remember the vertebrae-elongating-over-ontogeny stuff from Wedel et al. (2000b). Maybe that’s an argument for breaking up long, kitchen-sink papers into two or more separate publications?

This will be all too short, but I can’t let the publication of a new giant sauropod pass unremarked. Yesterday Bernardo González Riga and colleagues published a nice, detailed paper describing Notocolossus gonzalezparejasi, “Dr. Jorge González Parejas’s southern giant”, a new titanosaur from the Late Cretaceous of Mendoza Province, Argentina (González Riga et al. 2016). The paper is open access and freely available to the world.

As you can see from the skeletal recon, there’s not a ton of material known from Notocolossus, but among giant sauropods it’s actually not bad, being better represented than Argentinosaurus, Puertasaurus, Argyrosaurus, and Paralititan. In particular, one hindfoot is complete and articulated, and a good chunk of the paper and supplementary info are devoted to describing how weird it is.

But let’s not kid ourselves – you’re not here for feet, unless it’s to ask how many feet long this monster was. So how big was Notocolossus, really?

Well, it wasn’t the world’s largest sauropod. And to their credit, no-one on the team that described it has made any such superlative claims for the animal. Instead they describe it as, “one of the largest terrestrial vertebrates ever discovered”, and that’s perfectly accurate.

Any discussions of the size of Notocolossus will be driven by one of two elements: the humerus and the anterior dorsal vertebra. The humerus is 176 cm long, which is shorter than those of Giraffatitan (213 cm), Brachiosaurus(204 cm), and Turiasaurus (179 cm), but longer than those of Paralititan (169 cm), Dreadnoughtus(160 cm), and Futalognkosaurus (156 cm). Of course we don’t have a humerus for Argentinosaurus or Puertasaurus, but based on the 250-cm femur of Argentinosaurus, the humerus was probably somewhere around 200 cm. Hold that thought.

Top row: my attempt at a symmetrical Notocolossus dorsal, made by mirroring the left half of the fossil from the next row down. Second row: photos of the Notocolossus dorsal with missing bits outlined, from Gonzalez Riga et al (2016: fig. 2). Scale bar is 20 cm (in original). Third row: the only known dorsal vertebra of Puertasaurus, scaled to about the same size as the Notocolossus vertebra, from Novas et al. (2005: fig. 2).

The anterior dorsal tells a similar story, and this is where I have to give González Riga et al. some props for publishing such detailed sets of measurements in the their supplementary information. They Measured Their Damned Dinosaur. The dorsal has a preserved height of 75 cm – it’s missing the tip of the neural spine and would have been a few cm taller in life – and by measuring the one complete transverse process and doubling it, the authors estimate that when complete it would have been 150 cm wide. That is 59 inches, almost 5 feet. The only wider vertebra I know of is the anterior dorsal of Puertasaurus, at a staggering 168 cm wide (Novas et al. 2005). The Puertasaurus dorsal is also quite a bit taller dorsoventrally, at 106 cm, and it has a considerably larger centrum: 43 x 60 cm, compared to 34 x 43.5 cm for Notocolossus (anterior centrum diameters, height x width).

Centrum size is an interesting parameter. Because centra are so rarely circular, arguably the best way to compare across taxa would be to measure the max area (or, since centrum ends are also rarely flat, the max cross-sectional area). It’s late and this post is already too long, so I’m not going to do that now. But I have been keeping an informal list of the largest centrum diameters among sauropods – and, therefore, among all Terran life – and here they are (please let me know if I missed anyone):

(Fine print: I’m only logging each taxon once, by its largest vertebra, and I’m not counting the dorsoventrally squashed Giraffatitan cervicals which get up to 47 cm wide, and the “uncatalogued” Supersaurus dorsal is one I saw back in 2005 – it almost certainly has been catalogued in the interim.) Two things impress me about this list: first, it’s not all ‘exotic’ weirdos – look at the giant Oklahoma Apatosaurus hanging out halfway down the list. Second, Argentinosaurus and Puertasaurus pretty much destroy everyone else by a wide margin. Notocolossus doesn’t seem so impressive in this list, but it’s worth remembering that the “max” centrum diameter here is from one vertebra, which was likely not the largest in the series – then again, the same is true for Puertasaurus, Alamosaurus, and many others.

As for the estimated mass of Notocolossus, González Riga et al. (2016) did their due diligence. The sections on mass estimation in the main text and supplementary information are very well done – lucid, modest, and fair. Rather than try to summarize the good bit, I’ll just quote it. Here you go, from page 7 of the main text:

The [humeral] diaphysis is elliptical in cross-section, with its long axis oriented mediolaterally, and measures 770 mm in minimum circumference. Based on that figure, the consistent relationship between humeral and femoral shaft circumference in associated titanosaurian skeletons that preserve both of these dimensions permits an estimate of the circumference of the missing femur of UNCUYO-LD 301 at 936 mm (see Supplementary Information). (Note, however, that the dataset that is the source of this estimate does not include many gigantic titanosaurs, such as Argentinosaurus[5], Paralititan[16], and Puertasaurus[11], since no specimens that preserve an associated humerus and femur are known for these taxa.) In turn, using a scaling equation proposed by Campione and Evans[20], the combined circumferences of the Notocolossus stylopodial elements generate a mean estimated body mass of ~60.4 metric tons, which exceeds the ~59.3 and ~38.1 metric ton masses estimated for the giant titanosaurs Dreadnoughtus and Futalognkosaurus, respectively, using the same equation (see Supplementary Information). It is important to note, however, that subtracting the mean percent prediction error of this equation (25.6% of calculated mass[20]) yields a substantially lower estimate of ~44.9 metric tons for UNCUYO-LD 301. Furthermore, Bates et al.[21] recently used a volumetric method to propose a revised maximum mass of ~38.2 metric tons for Dreadnoughtus, which suggests that the Campione and Evans[20] equation may substantially overestimate the masses of large sauropods, particularly giant titanosaurs. Unfortunately, however, the incompleteness of the Notocolossus specimens prohibits the construction of a well-supported volumetric model of this taxon, and therefore precludes the application of the Bates et al.[21] method. The discrepancies in mass estimation produced by the Campione and Evans[20] and Bates et al.[21] methods indicate a need to compare the predictions of these methods across a broad range of terrestrial tetrapod taxa[21]. Nevertheless, even if the body mass of the Notocolossus holotype was closer to 40 than 60 metric tons, this, coupled with the linear dimensions of its skeletal elements, would still suggest that it represents one of the largest land animals yet discovered.

So, nice work all around. As always, I hope we get more of this critter someday, but until then, González Riga et al. (2016) have done a bang-up job describing the specimens they have. Both the paper and the supplementary information will reward a thorough read-through, and they’re free, so go have fun.

July 14, 2015

Now that the new Wilson and Allain (2015) paper has redescribed Rebbachisaurus, we can use it to start thinking about some other specimens. Particularly helpful is this beautiful rotating animation of the best dorsal vertebra (here captured at the point of the rotation where we’ve viewing it in right anterolateral):

Most obviously, there are laminae running up and down the anterior and posterior margins of the lateral face of Xeno’s neural arch, and those same laminae seem to exist in Rebbachisaurus. We didn’t name these laminae in the Xeno paper, but if they’re the same thing as in Rebbachisaurus, then they’re ACPLs and PCDL — anterior centroparapophyseal and posterior centodiapophyseal laminae.

If that’s right, then we misinterpreted the site of the parapophysis in Xenoposeidon. We (Taylor and Naish 2007) thought it was at the cross-shaped junction of laminae near the anterodorsalmost preserved part of the vertebra. In Rebbachisaurus, this cross exists, but it’s merely where the CPRL (centroprezygapophyseal lamina) intersects the ACPL.

But there’s more. In Xenoposeidon, the base of the CPRL (if that’s what it is) forms a “V” shape with an accessory lamina that proceeds posterodorsally from the same origin. (This is one of the features that’s apparent on the more damaged right side of Xeno as well as the nicer left side.) That lamina also seems to exist in Rebbachisaurus — but with the whole vertebra to consider, we can see that it’s not an accessory lamina, but a perfectly well-behaved CPOL (centropostzygapophseal lamina).

So if Xeno is indeed a rebbachisaurid, then the two branches of the “V” go to support the pre- and postzygs, and the laminae running up the anterior and posterior margins of the centrum support the parapophyses and diapophyses respectively. There are actually two crosses on each side of the neural arch: one at the intersection of CPRL and ACPL, the other at the intersection of CPOL and PCDL; but in the Xeno specimen, the posterior cross is lost, having been just above where the break occurs at the top of the neural arch.

Here’s what I mean:

In case it’s not clear, the grey lines are an (extremely crude) reconstruction, the blue lines label the important laminae, and the red circles highlight the two crosses.

Hmm. The more I look at this, the more convincing I find it.

But there’s more! The anterior aspect of the Rebbachisaurus vertebra also bears a notable resemblance to what we see in Xeno, with a pair or arched laminae forming a vaulted roof to the neural canal.

That’s not a bad call—the infrazygapophyseal region of that vert is elongate, and there is a nice CPRF and those closely positioned TPRLs could mean that prz’s are close to one another or even conjoined. It’s tantalizing, but not much to go on. Would be nice to prep out CPRF & work out laminae on lat sfc.

Jeff is right that more preparation would help to figure this out.

Not that everything about the Rebbachisaurus dorsal is Xeno-ish. Most notably, the lateral foramen is nothing like that of Xeno, being an uninspiringly dull and simple oval rather than the much more elegant foramen-within-a-fossa arrangement that we see in Xeno. But there are other points of commonality, too, such as the flat stretch of bone above the fossa and the way the posterior margin of the neural arch reaches the posterior margin of the centrum.

All in all — while there is plenty of work yet to do — I am increasingly inclined to think that the evidence we currently have suggests Xenoposeidon is a rebbachisaurid. If that’s right, it would be quite an exciting result. It would be the earliest known rebbachisaur, and the only named one from the UK. (Mannion 2009 described, but did not name, a rebbachisaurid scapula from Wessex formation of the Wealden). Could Mannion’s scapula be Xenoposeidon? Unlikely, as it’s 10 million years more recent. But it could be a close relative.

Exciting times!

References

Mannion, Philip D. 2009. A rebbachisaurid sauropod from the Lower Cretaceous of the Isle of Wight, England. Cretaceous Research30:521-526. doi:10.1016/j.cretres.2008.09.005

July 13, 2015

Here at SV-POW! Towers, we’re keenly aware that some of our fans are just here for the hardcore sauropod vertebra action. These folks start to shift in their seats when we put up too many posts in a row on open access or rabbits or…okay, mostly just OA and bunnies. If that’s you – or, heck, even if it isn’t – your good day has come. Saddle up. Let’s ride.

When Brian Engh and I were at the new Natural History Museum of Utah recently, I spotted this cute little juvenile cervical in one of the display cases.

According to this sign, it’s UMNH 21054, and it was found by Frank DeCourten and prepared by Virginia Tidwell.

It shares a display case and a sign with what is probably an anterior dorsal, UMNH 21055.

Now, I don’t mean to brag (okay, maybe a little…) but the number of EKNApod* vertebrae is not large and the number of EKNApod vertebrae I’m not intimately familiar with hovers near zero. This thing was ringing bells – I knew I’d seen it before.

* Early Cretaceous North American sauropod

Here are few more views. Note the light-colored oblong spot on the top of the condyle in the image above – this may be a pneumatic foramen filled with matrix, or a spot where the cortical bone flaked away to reveal one of the internal pneumatic spaces. Also, check out the fragment of extraneous bone (probably cervical rib) stuck sideways across the top of the centrum, just behind the condyle, in the image immediately below. Both of these features will be important later.

The vert belongs to a juvenile sauropod because the neural arch is missing – it didn’t fuse to the centrum before the animal died. But it was a big baby; the centrum is maybe just a hair under 40 cm in length, meaning that a world-record giraffe might just maybe have a couple of cervicals of the same length. But basal titanosauriforms typically have 12-13 cervicals, not the whimpy 7 that almost all mammals must make do with, and all-stars like Euhelopus can have up to 17.

Also, this was not from the middle of the neck. No way. The parapophyses are huge, and the centrum is pretty stubby compared to Sauroposeidon or YPM 5294, the Sauroposeidonesque cervical from Unit VII of the Cloverly (pic here). My guess is we’re looking at something past the middle of the neck, where the cervicals start to get proportionally shorter (but sometimes max out in absolute length), maybe a C9 or C10. In Giraffatitan brancai HM SII/MB.R.2181, C10 has a centrum length of 100 cm and makes up about 12% of the 8.5-meter neck. Assuming similar proportions here, UMNH 21054 came from the roughly 3-meter neck of a sauropod about the size of a really big draft horse or a really small elephant.

But enough noodling about the animal’s size. I knew I’d seen this vert before, but where? Thank goodness for comprehensive signage – I knew the material had been discovered by Frank DeCourten and prepped by Virginia Tidwell. At one of the SVP meetings in Denver, at a reception at the Denver museum, Virginia had invited me into the prep lab to see some EKNApod material from the Long Walk Quarry in Utah. The Long Walk Quarry was Frank DeCourten’s baby – he wrote a couple of papers about it (e.g., DeCourten 1991) and included additional information in his book, Dinosaurs of Utah (1998; second edition in 2013). DeCourten had referred the material to Pleurocoelus because that’s what people did with EKNApods back in the 20th century, but I remembered seeing one cervical that, like Sauroposeidon and YPM 5294, was just too long to match any of the Pleurocoelus material. My ‘Museum Photos’ file has a subfolder titled ‘Denver 2004’ – was the mystery vert in there?

In short, yes. Here’s one of the photos I took back in 1994.

Here’s another, sans flash this time. Check out the white spot on top of the condyle, the bar of float bone stuck sideways across the centrum just behind the spot, and general pattern of breaks – it’s a perfect match for UMNH 21054. Also note the block number on the pink specimen label at the bottom of the image – LWQ8, for Long Walk Quarry.

Three mysteries remain. One, the signage says the vert is from Carbon County, Utah, but the Long Walk Quarry has always been described as being in Emery County. Just a typo, or is there a story there? Two, how much of the animal (or animals) was excavated and prepped? I saw other vertebrae, both larger and smaller, when I was in Denver back on ’04, and DeCourten figured still others that I haven’t yet seen personally. Finally, is anyone working on it? And if not…[cautiously raises hand].

Right on the heels of Aquilopslast week, my paper with John Foster on the new specimen of Haplocanthosaurus from Snowmass, Colorado, was just published in Volumina Jurassica. I’ll have more to say about it later, but right now I am up against a deadline on a big project and I need to go work on that. I’m only popping up here to note two quick things.

First, if you’re not familiar with Volumina Jurassica – and I wasn’t, before this project – it’s a free-to-access* journal that publishes papers on all aspects of the Jurassic. The current issue is specifically dedicated to the Jurassic formations of the American West. There’s a lot of interesting stuff in there, but of special interest to SV-POW! readers will be the paper by Cary Woodruff and John Foster on the legendary and possibly apocryphal Amphicoelias fragillimus.

Fig. 4. Sacra of Haplocanthosaurus. A. MWC 8028, sacrum in right lateral view. B. MWC 8028, close-up of S4 and S5 centra highlighting pneumatic fossae. C. MWC 8028 with divisions between the vertebrae overlaid. D. CM 879, sacrum in right lateral view with divisions between the vertebrae overlaid. E. CM 572 in right lateral view, after Hatcher (1903c: plate 4). B–E are not shown at the same scale, scale bar for A = 20 cm. Note that the neural arches in CM 572 were restored during preparation, and the sacral neural spines as shown here are probably lower than they would have been in life.

Second, the figure resolution in the PDF of the Haplocanthosaurus paper is not stellar, so as is the case with almost all of our papers, the full-color, high-resolution figures are available at the paper’s page on the sidebar.

July 16, 2014

I was at the Natural History Museum of Los Angeles County yesterday to do some research in the ornithology collection. After lunch I was working on this pelican skeleton and I thought, “Geez, there is just no way to do this thing justice with still photos. I should make a video.” Here it is. You’ll want to see it full-screen–this being my first time out making a video, I didn’t realize that I was holding the phone the wrong way for efficient viewing on other devices.

The specimen is LACM Ornithology 86262. I’m posting this video with the knowledge and kind permission of the ornithology collection staff.